Previous measurements of mammalian photoreceptor spectral sensitivity have been analysed, with particular attention to the long-wavelength region. The measurements selected for study come from rod and cone systems, and from human, monkey, bovine and squirrel sources. For the spectra from photoreceptor electrophysiology and from psychophysical sensitivity, the frequency scaling applied by Mansfield (1985, The visual system, pp. 89-106. New York: Alan Liss) provides a common shape over a range of at least 7 log10 units of sensitivity, from low frequencies (long wavelengths) to frequencies beyond the peak. The same curve is applicable to the absorbance spectrum of bovine rhodopsin, although the absorbance can only be measured down to about 2 log10 units below the peak. At the longest wavelengths the results exhibit a common limiting slope of 70 loge units (or 30.4 log10 units) per unit of normalized frequency. A simple equation is presented as a generic description for the alpha-band of mammalian photoreceptor spectral sensitivity curves, and it seem likely that the equation may be equally applicable to retinal1-based pigments in other species. Despite the lack of a theoretical basis, the equation has the correct asymptotic behaviour at long wavelengths, and it provides an accurate description of the peak. It also accounts accurately for the experimentally observed "yellowing" of long-wavelength lights that occurs beyond 700 nm.

We examined the responses of toad rod photoreceptors to single photons of light. To minimize the effects of variability in the early rising phase, we selected sets of responses that closely matched the rise of the mean single photon response. Responses selected in this way showed substantial variations in kinetics, appearing to peel off from a common time course after different delays. Following incorporation of the calcium buffer BAPTA, the time to peeling off was retarded. Our analysis indicates that it is not necessary to invoke a long series of reaction steps to explain the shutoff of rhodopsin activity. Instead, our results suggest that the observed behavior is explicable by the presently known shutoff reactions of activated rhodopsin, modulated by feedback.

A minimally invasive technique is described for recording the a-wave of the human ERG and extracting the parameters of transduction in the rod and cone photoreceptors. A corneal DTL fibre electrode is used, but the pupil is not dilated and the cornea is not anaesthetized. Although the amplitude of the signal collected by the DTL electrode varies from session to session, this is not a problem, as the photoreceptor fractional circulating current is obtained by normalization of the response family. A method is described for varying the effective flash intensity over a wide range, by controlling the duration of the xenon flash. In order to fit the kinetics of the responses, an analytical equation is derived for the convolution of the previous "delayed gaussian" expression with the cell's capacitive time constant. This equation provides a good description of both the rod and the cone response families. For rods, the capacitive time constant was found to be tau rod approximately 1 msec as reported previously, but for the cones a considerably longer time constant of tau cone approximately 4-5 msec was needed. For rods, the amplification constant (Arod approximately 5 sec-2) was close to previous estimates, but for cones the sensitivity (expressed in terms of corneal illuminance) was higher than in previous work. Calculation of the amplification constant of transduction within the cones requires knowledge of their light collection properties, and the absence of hard information makes this estimate somewhat speculative. However, when account is taken of the larger diameter of the inner segments of cones in the peripheral retina, then our estimated amplification constant for the cones (Acone approximately 3-7 sec-2) is of a similar order of magnitude to that obtained for the rods.

Both cGMP and Ca2' appear to be involved in the process of phototransduction in vertebrate rods, but their precise roles have been the subject of debate. To investigate the role of Ca21 we have artificially increased the calcium buffering capacity of the rod by using a patch pipet to incorporate the calcium buffer 1,2-bis(o-aminophenoxy)ethane-N,N,N',N'-tetraacetic acid (BAPTA) into the rod cytoplasm. In the presence of buffer the Na'-Ca2' exchange current became greatly slowed, suggesting that the cytoplasmic calcium concentration (Cai) had indeed been buffered substantially. Although the presence of buffer had negligible effect on the rising phase of the light response, it profoundly altered the later behavior. Responses to brief flashes became prolonged and exhibited an overshoot, apparently because the shut-off process was modified. The normal acceleration of time-to-peak with brighter flashes (an early sign of light adaptation) disappeared. Responses to steady adapting illumination took much longer than normal to settle to a steady level, although the final level represented a similar fractional suppression of current. With superimposed test flashes the presence of such adapting illumination caused a more rapid recovery, whereas the presence of calcium buffer slowed the recovery. The results are consistent with the idea that the rapid drop in Ca1, which has recently been shown to accompany the light response, is involved in terminating the light response, and that Cai is thereby involved in setting the operating point and sensitivity of phototransduction. From comparison with other work we infer that Cai appears to act, at least in part, by means of control of cGMP phosphodiesterase activity.The electrical response of vertebrate photoreceptors to illumination consists of a hyperpolarization (1, 2) brought about by the closure of ion channels in the outer segment plasma membrane (2, 3). The transduction process has very high amplification under dark-adapted conditions, and the absorption of a photon by a single rhodopsin molecule can block the entry into the outer segment of more than 106 Na+ ions (4). In the presence of background illumination the photoreceptor becomes light-adapted, and the gain of transduction decreases in approximately inverse proportion to the light intensity (5, 6), in cones and in rods of lower vertebrates. This automatic gain control is probably of great importance to the operation of the visual system.The need for a diffusible internal messenger substance mediating transduction was recognized from the outset (2, 7), and the proposal was originally made that channel closure was brought about by the release of Ca2+ into the cytoplasm (7). Subsequently the importance of cGMP was recognized, and a cascade of biochemical reactions leading to the activation of a cGMP phosphodiesterase (PDEase) was described; many groups were involved in these developments and the subject is reviewed in refs. 8 and 9. For some years, however, it was not clear whether either or both Ca2l and cGMP we...

Sensory transduction shares common features in widely different sensory modalities. The purpose of this article is to examine the similarities and differences in the underlying mechanisms of transduction in the sensory receptor cells for vision, olfaction, and hearing. One of the major differences between the systems relates to the nature of the stimulus. In both the visual and olfactory systems a quantal mechanism of detection is possible, because the absorption of a photon or the binding of an odorant molecule provides an energy change significantly greater than the thermal noise in the receptor molecule. In hearing, on the other hand, the energy of a phonon is far lower, and detection occurs by a "classical" mechanism. For vertebrate photoreceptors and olfactory receptor cells, sensory transduction employs a G protein cascade that is remarkably similar in the two cases, and that is closely homologous to other G protein signaling cascades. For auditory and vestibular hair cells, transduction operates via a mechanism of direct coupling of the stimulus to ion channels, in a manner reminiscent of the direct gating of post-synaptic ion channels in various synaptic mechanisms. The three classes of sensory receptor cell share similarities in their mechanisms of adaptation, and it appears in each case that cytoplasmic calcium concentration plays a major role in adaptation.